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Modeling Sexual Differences of Body Size Variation in Ground Beetles In Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 20 October 2020 doi:10.20944/preprints202010.0408.v1 Article Modeling Sexual Differences of Body Size Variation in Ground Beetles in Geographical Gradient (the case study in Pterostichus oblongopunctatus Fabricius, 1787) Raisa Sukhodolskaya 1, Anatoliy Saveliev 2, Nadezhda Ukhova 3, Iraida. Vorobyova 4, Igor Solodovnikov 5, Anatoliy Anciferov 6, Tatyana Gordienko 1, Rifgat Shagidullin 1 and Dmitriy Vavilov 1 1 The Institute of Problems in Ecology and Mineral Wealth, Tatarstan Academy of Sciences; [email protected] 2 Kazan (Volga Region) Federal University; [email protected] 3 Visimskiy Nature Reserve; [email protected] 4 Mariy State University; [email protected] 5 Educational establishment «Vitebsk State P.M. Masherov University; Igor A. Solodovnikov 6 Kostroma Museum-Reserve; [email protected] * Correspondence: [email protected]; Tel.: +79503152619 Simple Summary: Body sizes of organisms vary considerably among species and among populations within species Geographic variation in resource availability and interspecific or intraspecific competition are likely important sources of variation in selection on body size of many animals, especially those with limited dispersal capacity. We investigated six traits in one of the ground beetle species in geographical gradient including three degrees in latitude and twenty nine degrees in longitude. We registered saw-tooth pattern. Females were always larger than males. Abstract: Concretizing the mechanisms of Bergmann rule, we found saw-tooth pattern in body size variation in ground beetle Pterostichus oblongopunctatus. We sampled beetles in 2010 – 2018 at the forest undisturbed plots on the broad territory in Russia. Investigating regions covered territory, extending to 3 degrees latitude and 19 degrees longitude. We measured six traits in every of 3294 caught individual. ANOVA showed that latitude, and sex affected significantly body size of the species studied. Mean values of each trait changed significantly from one studied region to another in females and males as well. Sexual size dimorphism in species was female-biased. We performed models in R to estimate the steepness of body size variation in both sexes. In overwhelming majority of cases that parameter was equal in both sexes. So the hypothesis, that male′s variation is steeper in latitude gradient was not confirmed. Keywords: body size variation; sexual size dimorphism; ground beetles; geographical gradient; modeling; saw-tooth pattern; Bergmann rule 1. Introduction Large-scale patterns of body size variation have occupied biologists for over a century. One prominent pattern is Bergmann’s rule, which originally referred to clinal geographic variation among endothermic (warm-blooded) species only. The latter tend to be bigger in colder climates [1-4]. That phenomena was explained as follows: larger individuals possess smaller surface-to-volume ratios and it is more conducive to conserving heat in cold climates. Then this rule was extended to ectothermic (cold-blooded) organisms [5]. For them the cause must be different, as especially small ectotherms acclimate to ambient temperature almost instantly [6]. © 2020 by the author(s). Distributed under a Creative Commons CC BY license. Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 20 October 2020 doi:10.20944/preprints202010.0408.v1 Among large-scale patterns of body size variation counter gradient variation [7] and saw-tooth variation exist. The latter considers, that variation in number of generations per season (i.e. voltinism) affects time constraints. The traditional view about how latitudinal or altitudinal gradient in season length affects the evolution of development time and body size in potentially multivoltine insects was developed by S. Masaki [8,9] and formalized by D. Roff [10,11] and Y. Iwasa et al. [12]. They assumed that development time and body size are positively correlated among genotypes, and insects use all available time for growth to maximize adult body size and thus fecundity. Consequently, both development time and body size are expected to increase along a gradient of increasing season length (i.e. with decreasing latitude or altitude) as long as voltinism does not change. Decreases in both traits are expected at the season lengths that facilitate the emergence of a new generation within the season, because time available per generation decreases at these transitions. Thus, the traditional saw-tooth cline arises across phenologies because of genetic adaptation to local conditions. The examples of saw-tooth variation were presented in recent years [13,14]. In addition the third pattern of body size variation in latitude gradient can occur. It is counter- gradient variation, known as the hypothesis of latitude compensation [15]. The mechanism can be described as follows: in high latitudes specimen compensate short breeding period by higher (it is genetically coded) rate of growth, compared with conspecificians in low latitudes. In simplest case the rate of growth is the body size per the unit of development time, the utterly exact compensation of breeding season constraints at the certain latitude (the environmental component) will occur in that situation, when growth rate (genetic component) evolves in the way to keep phenotypic body size equal at all latitudes. If increasing growth rate perfectly compensates for the decrease in generation time, body size remains constant over the whole gradient of season length. No less important aspect in studying Bergmann′s clines is the difference of their occurrence in males and females. Prominent body size pattern is Rensch’s rule, the observation widespread in the animal kingdom that male body size varies (or evolutionarily diverges) more than female body size among related species, such that male-biased sexual size dimorphism increases and female-biased sexual size dimorphism decreases with body size [16-18]. Rensch’s rule is manifested statistically in allometric slopes greater than one when male size is regressed on female size [18]. The problem regarding the scaling of SSD with body size exists. In the first place, there is the question of the taxonomic level at which it is studied – inter-specific or intra-specific. Most studies of the scaling of SSD with body size either phylogenetically- based or not have been performed across species at different levels (Fairbairn et al., 2007). An additional problem is that of the appropriate measurements for analyzing SSD (Fairbairn, 2007). Taking into account that different traits variation can be different in the same species, preferably several traits must be under investigation (Bidau et al., 2016). The few studies to date testing whether intraspecific variation in sexual size dimorphism conforms to Rensch’s rule have yielded mixed results [22-26]. Blanckenhorn et al. [27] in the large review studied latitude clines in males and females taking into account published data and their own investigations. Males latitudinal clines were steeper than those of females in a significant majority of species (66 of 98), confirming the male - steeper hypothesis connecting Bergmann’s and Rensch’s rules, regardless of whether the species displays male- or female-biased size dimorphism. That supported the existence of a geographic version of Rensch’s rule. Of course, that left approximately one-third of the species showing the opposite trend, underscoring the fact that several factors must be involved in generating those patterns. As previously shown for arthropods [28] they found a continuum of Bergmann (58 species) and converse Bergmann clines (40 species) in this taxonomically more diverse data set [29]. Interestingly, regardless of which sex was larger, the latitudinal decrease in male size (relative to female size), and hence change in sexual size dimorphism, in species with converse Bergmann clines was generally more pronounced than the corresponding relative male size increase with latitude in species with Bergmann clines, implying that dimorphism changed faster with latitude in the former group. The authors concluded that relationship between cline type and magnitude of divergence in male and female size required further study. Thus and so, we tried to replenish the cited above study with new research improving data analysis design. We investigated latitudinal body size variation in carabid beetle Pterostichus Preprints (www.preprints.org) | NOT PEER-REVIEWED | Posted: 20 October 2020 doi:10.20944/preprints202010.0408.v1 oblongopunctatus Fabricius, 1787. Ground beetles are famous for their indicative properties because of their high sensitivity to environmental effects [30]. This is a large species-rich family in Coleoptera. Despite plenty of papers on carabids, very few discuss their body size variation along geographical gradients. Among them K. Homburg investigation, revealing hump-shaped body size variation in North Africa – North Europe region [31], but that work was done on interspecific level. Intra-specific studies in carabids body size variation in latitude/altitude gradient are scarce. They are often limited by the single trait investigating or are done on the community level [32-35]. Ground beetles vary greatly in size and shape [36] but not in the magnitude of SSD [37]. Some species are fairly common. They are easy to collect in a large range of biotopes and they have large geographic distributions. These allows the sampling of several populations
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